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1
Singchat, Worapong, Syed Farhan Ahmad, Nararat Laopichienpong, Aorarat Suntronpong, Thitipong Panthum, Darren K. Griffin, and Kornsorn Srikulnath. "Snake W Sex Chromosome: The Shadow of Ancestral Amniote Super-Sex Chromosome." Cells 9, no. 11 (October 31, 2020): 2386. http://dx.doi.org/10.3390/cells9112386.
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Heteromorphic sex chromosomes, particularly the ZZ/ZW sex chromosome system of birds and some reptiles, undergo evolutionary dynamics distinct from those of autosomes. The W sex chromosome is a unique karyological member of this heteromorphic pair, which has been extensively studied in snakes to explore the origin, evolution, and genetic diversity of amniote sex chromosomes. The snake W sex chromosome offers a fascinating model system to elucidate ancestral trajectories that have resulted in genetic divergence of amniote sex chromosomes. Although the principal mechanism driving evolution of the amniote sex chromosome remains obscure, an emerging hypothesis, supported by studies of W sex chromosomes of squamate reptiles and snakes, suggests that sex chromosomes share varied genomic blocks across several amniote lineages. This implies the possible split of an ancestral super-sex chromosome via chromosomal rearrangements. We review the major findings pertaining to sex chromosomal profiles in amniotes and discuss the evolution of an ancestral super-sex chromosome by collating recent evidence sourced mainly from the snake W sex chromosome analysis. We highlight the role of repeat-mediated sex chromosome conformation and present a genomic landscape of snake Z and W chromosomes, which reveals the relative abundance of major repeats, and identifies the expansion of certain transposable elements. The latest revolution in chromosomics, i.e., complete telomere-to-telomere assembly, offers mechanistic insights into the evolutionary origin of sex chromosomes.
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Marec, František, and Walther Traut. "Sex chromosome pairing and sex chromatin bodies in W–Z translocation strains of Ephestia kuehniella (Lepidoptera)." Genome 37, no. 3 (June 1, 1994): 426–35. http://dx.doi.org/10.1139/g94-060.
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Structure and pairing behavior of sex chromosomes in females of four T(W;Z) lines of the Mediterranean flour moth, Ephestia kuehniella, were investigated using light and electron microscopic techniques and compared with the wild type. In light microscopic preparations of pachytene oocytes of wild-type females, the WZ bivalent stands out by its heterochromatic W chromosome strand. In T(W;Z) females, the part of the Z chromosome that was translated onto the W chromosome was demonstrated as a distal segment of the neo-W chromosome, displaying a characteristic non-W chromosomal chromomere–interchromomere pattern. This segment is homologously paired with the corresponding part of a complete Z chromosome. In contrast with the single ball of heterochromatic W chromatin in highly polyploid somatic nuclei of wild-type females, the translocation causes the formation of deformed or fragmented W chromatin bodies, probably owing to opposing tendencies of the Z and W chromosomal parts of the neo-W. In electron microscopic preparations of microspread nuclei, sex chromosome bivalents were identified by the remnants of electron-dense heterochromatin tangles decorating the W chromosome axis, by the different lengths of the Z and W chromosome axes, and by incomplete pairing. No heterochromatin tangles were attached to the translocated segment of the Z chromosome at one end of the neo-W chromosome. Because of the homologous pairing between the translocation and the structurally normal Z chromosome, pairing affinity of sex chromosomes in T(W;Z) females is significantly improved. Specific differences observed among T(W;Z)1–4 translocations are probably due to the different lengths of the translocated segments.Key words: Mediterranean flour moth, sex chromosomes, sex chromatin, translocations, synaptonemal complexes, microspreading.
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Uno, Yoshinobu, Chizuko Nishida, Chiyo Takagi, Takeshi Igawa, Naoto Ueno, Masayuki Sumida, and Yoichi Matsuda. "Extraordinary Diversity in the Origins of Sex Chromosomes in Anurans Inferred from Comparative Gene Mapping." Cytogenetic and Genome Research 145, no. 3-4 (2015): 218–29. http://dx.doi.org/10.1159/000431211.
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Sex determination in frogs (anurans) is genetic and includes both male and female heterogamety. However, the origins of the sex chromosomes and their differentiation processes are poorly known. To investigate diversity in the origins of anuran sex chromosomes, we compared the chromosomal locations of sex-linked genes in 4 species: the African clawed frog (Xenopus laevis), the Western clawed frog (Silurana/X. tropicalis), the Japanese bell-ring frog (Buergeria buergeri), and the Japanese wrinkled frog (Rana rugosa). Comparative mapping data revealed that the sex chromosomes of X. laevis, X. tropicalis and R. rugosa are different chromosome pairs; however, the sex chromosomes of X. tropicalis and B. buergeri are homologous, although this may represent distinct evolutionary origins. We also examined the status of sex chromosomal differentiation in B. buergeri, which possesses heteromorphic ZW sex chromosomes, using comparative genomic hybridization and chromosome painting with DNA probes from the microdissected W chromosome. At least 3 rearrangement events have occurred in the proto-W chromosome: deletion of the nucleolus organizer region and a paracentric inversion followed by amplification of non-W-specific repetitive sequences.
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Hejníčková, Martina, Martina Dalíková, Pavel Potocký, Toomas Tammaru, Marharyta Trehubenko, Svatava Kubíčková, František Marec, and Magda Zrzavá. "Degenerated, Undifferentiated, Rearranged, Lost: High Variability of Sex Chromosomes in Geometridae (Lepidoptera) Identified by Sex Chromatin." Cells 10, no. 9 (August 28, 2021): 2230. http://dx.doi.org/10.3390/cells10092230.
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Sex chromatin is a conspicuous body that occurs in polyploid nuclei of most lepidopteran females and consists of numerous copies of the W sex chromosome. It is also a cytogenetic tool used to rapidly assess the W chromosome presence in Lepidoptera. However, certain chromosomal features could disrupt the formation of sex chromatin and lead to the false conclusion that the W chromosome is absent in the respective species. Here we tested the sex chromatin presence in 50 species of Geometridae. In eight selected species with either missing, atypical, or normal sex chromatin patterns, we performed a detailed karyotype analysis by means of comparative genomic hybridization (CGH) and fluorescence in situ hybridization (FISH). The results showed a high diversity of W chromosomes and clarified the reasons for atypical sex chromatin, including the absence or poor differentiation of W, rearrangements leading to the neo-W emergence, possible association with the nucleolus, and the existence of multiple W chromosomes. In two species, we detected intraspecific variability in the sex chromatin status and sex chromosome constitution. We show that the sex chromatin is not a sufficient marker of the W chromosome presence, but it may be an excellent tool to pinpoint species with atypical sex chromosomes.
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Rutkowska, Joanna, Malgorzata Lagisz, and Shinichi Nakagawa. "The long and the short of avian W chromosomes: no evidence for gradual W shortening." Biology Letters 8, no. 4 (March 14, 2012): 636–38. http://dx.doi.org/10.1098/rsbl.2012.0083.
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The well-established view of the evolution of sex chromosome dimorphism is of a gradual genetic and morphological degeneration of the hemizygous chromosome. Yet, no large-scale comparative analysis exists to support this view. Here, we analysed karyotypes of 200 bird species to test whether the supposed directional changes occur in bird sex chromosomes. We found no support for the view that W chromosomes gradually become smaller over evolutionary time. On the contrary, the length of the W chromosome can fluctuate over short time scales, probably involving both shortening and elongation of non-coding regions. Recent discoveries of near-identical palindromes and neo-sex chromosomes in birds may also contribute to the observed variation. Further studies are now needed to investigate how chromosome morphology relates to its gene content, and whether the changes in size were driven by selection.
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Lisachov, Artem P., Svetlana A. Galkina, Alsu F. Saifitdinova, Svetlana A. Romanenko, Daria A. Andreyushkova, Vladimir A. Trifonov, and Pavel M. Borodin. "Identification of sex chromosomes in Eremias velox (Lacertidae, Reptilia) using lampbrush chromosome analysis." Comparative Cytogenetics 13, no. 2 (May 14, 2019): 17–28. http://dx.doi.org/10.3897/compcytogen.v13i2.34116.
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Reptiles are good objects for studying the evolution of sex determination, since they have different sex determination systems in different lineages. Lacertid lizards have been long-known for possessing ZZ/ZW type sex chromosomes. However, due to morphological uniformity of lacertid chromosomes, the Z chromosome has been only putatively cytologically identified. We used lampbrush chromosome (LBC) analysis and FISH with a W-specific probe in Eremiasvelox (Pallas, 1771) to unequivocally identify the ZW bivalent and investigate its meiotic behavior. The heterochromatic W chromosome is decondensed at the lampbrush stage, indicating active transcription, contrast with the highly condensed condition of the lampbrush W chromosomes in birds. We identified the Z chromosome by its chiasmatic association with the W chromosome as chromosome XIII of the 19 chromosomes in the LBC karyotype. Our findings agree with previous genetic and genomic studies, which suggested that the lacertid Z chromosome should be one of the smaller macrochromosomes.
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Mahony, M. J. "Heteromorphic sex chromosomes in the Australian frog Crinia bilingua (Anura: Myobatrachidae)." Genome 34, no. 3 (June 1, 1991): 334–37. http://dx.doi.org/10.1139/g91-055.
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The karyotype of Crinia bilingua was examined and analysed with standard staining, C-banding, and silver-staining. Heteromorphic sex chromosomes of the ZW ♂/ZZ ♀ type were observed. The larger W chromosome is submetacentric and the smaller Z chromosome is acrocentric. The centromere and proximal region of the short arm of the W chromosome consist of constitutive heterochromatin (C-band region), and beyond this is a small euchromatic terminal region. The centromere of the Z chromosome did not C-band. The long arms of the Z and W chromosomes are euchromatic and equal in length. The nucleolar organiser region occurs terminally on the long arm of both the Z and W chromosomes, and there is no cytological evidence for inactivity of the nucleolar organiser region on the W chromosome. These features indicate an early stage in the evolution of heteromorphic sex chromosomes.Key words: heteromorphic sex chromosomes, frog, Crinia bilingua.
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Scacchetti, Priscilla C., Ricardo Utsunomia, José C. Pansonato-Alves, Marcelo R. Vicari, Roberto F. Artoni, Claudio Oliveira, and Fausto Foresti. "Chromosomal Mapping of Repetitive DNAs in Characidium (Teleostei, Characiformes): Genomic Organization and Diversification of ZW Sex Chromosomes." Cytogenetic and Genome Research 146, no. 2 (2015): 136–43. http://dx.doi.org/10.1159/000437165.
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The speciose neotropical genus Characidium has proven to be a good model for cytogenetic exploration. Representatives of this genus often have a conserved diploid chromosome number; some species exhibit a highly differentiated ZZ/ZW sex chromosome system, while others do not show any sex-related chromosome heteromorphism. In this study, chromosome painting using a W-specific probe and comparative chromosome mapping of repetitive sequences, including ribosomal clusters and 4 microsatellite motifs - (CA)15, (GA)15, (CG)15, and (TTA)10 -, were performed in 6 Characidium species, 5 of which possessed a heteromorphic ZW sex chromosome system. The W-specific probe showed hybridization signals on the W chromosome of all analyzed species, indicating homology among the W chromosomes. Remarkably, a single major rDNA-bearing chromosome pair was found in all species. The 18S rDNA localized to the sex chromosomes in C. lanei, C. timbuiense and C. pterostictum, while the major rDNA localized to one autosome pair in C. vidali and C. gomesi. In contrast, the number of 5S rDNA-bearing chromosomes varied. Notably, minor ribosomal clusters were identified in the W chromosome of C. vidali. Microsatellites were widely distributed across almost all chromosomes of the karyotypes, with a greater accumulation in the subtelomeric regions. However, clear differences in the abundance of each motif were detected in each species. In addition, the Z and W chromosomes showed the differential accumulation of distinct motifs. Our results revealed variability in the distribution of repetitive DNA sequences and their possible association with sex chromosome diversification in Characidium species.
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Sharbel, Timothy F., David M. Green, and Andreas Houben. "B-chromosome origin in the endemic New Zealand frog Leiopelma hochstetteri through sex chromosome devolution." Genome 41, no. 1 (February 1, 1998): 14–22. http://dx.doi.org/10.1139/g97-091.
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The endemic New Zealand frog Leiopelma hochstetteri has variable numbers of mitotically stable B chromosomes. To assess whether the B chromosomes were derived from the autosome complement, they were isolated by micromanipulation and their DNA amplified by degenerate oligonucleotide primed PCR. Southern hybridizations of B chromosome DNA probes to genomic DNA from males and females characterized by differing numbers of B chromosomes demonstrated that the B chromosomes were derived from the univalent W sex chromosome characteristic of North Island populations. The presence of homologous B chromosome specific sequences from geographically distinct populations indicates a single origin of the B chromosomes. Furthermore, a primitive homology shared by B chromosomes and the W sex chromosome from an ancestral WZ/ZZ karyotype, which is still present in frogs from Great Barrier Island, shows that the B chromosomes originated soon after the univalent W sex chromosome had originated. Sequence analysis revealed that B chromosome DNA is composed of repeat sequences and has the potential to form stable hairpin structures. The molecular dynamics of these structures may reflect an inherent propensity to undergo rapid change in nucleotide sequence and chromosome structure.
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Johnson Pokorná, Martina, Marie Altmanová, Michail Rovatsos, Petr Velenský, Roman Vodička, Ivan Rehák, and Lukáš Kratochvíl. "First Description of the Karyotype and Sex Chromosomes in the Komodo Dragon (Varanus komodoensis)." Cytogenetic and Genome Research 148, no. 4 (2016): 284–91. http://dx.doi.org/10.1159/000447340.
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The Komodo dragon (Varanus komodoensis) is the largest lizard in the world. Surprisingly, it has not yet been cytogenetically examined. Here, we present the very first description of its karyotype and sex chromosomes. The karyotype consists of 2n = 40 chromosomes, 16 macrochromosomes and 24 microchromosomes. Although the chromosome number is constant for all species of monitor lizards (family Varanidae) with the currently reported karyotype, variability in the morphology of the macrochromosomes has been previously documented within the group. We uncovered highly differentiated ZZ/ZW sex microchromosomes with a heterochromatic W chromosome in the Komodo dragon. Sex chromosomes have so far only been described in a few species of varanids including V. varius, the sister species to Komodo dragon, whose W chromosome is notably larger than that of the Komodo dragon. Accumulations of several microsatellite sequences in the W chromosome have recently been detected in 3 species of monitor lizards; however, these accumulations are absent from the W chromosome of the Komodo dragon. In conclusion, although varanids are rather conservative in karyotypes, their W chromosomes exhibit substantial variability at the sequence level, adding further evidence that degenerated sex chromosomes may represent the most dynamic genome part.
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Pucci, Marcela B., Patricia Barbosa, Viviane Nogaroto, Mara C. Almeida, Roberto F. Artoni, Priscila C. Scacchetti, José C. Pansonato-Alves, Fausto Foresti, Orlando Moreira-Filho, and Marcelo R. Vicari. "Chromosomal Spreading of Microsatellites and (TTAGGG)n Sequences in the Characidium zebra and C. gomesi Genomes (Characiformes: Crenuchidae)." Cytogenetic and Genome Research 149, no. 3 (2016): 182–90. http://dx.doi.org/10.1159/000447959.
Full textAbstract:
Sex chromosome evolution involves the accumulation of repeat sequences such as multigenic families, noncoding repetitive DNA (satellite, minisatellite, and microsatellite), and mobile elements such as transposons and retrotransposons. Most species of Characidium exhibit heteromorphic ZZ/ZW sex chromosomes; the W is characterized by an intense accumulation of repetitive DNA including dispersed satellite DNA sequences and transposable elements. The aim of this study was to analyze the distribution pattern of 18 different tandem repeats, including (GATA)n and (TTAGGG)n, in the genomes of C. zebra and C. gomesi, especially in the C. gomesi W chromosome. In the C. gomesi W chromosome, weak signals were seen for (CAA)10, (CAC)10, (CAT)10, (CGG)10, (GAC)10, and (CA)15 probes. (GA)15 and (TA)15 hybridized to the autosomes but not to the W chromosome. The (GATA)n probe hybridized to the short arms of the W chromosome as well as the (CG)15 probe. The (GATA)n repeat is known to be a protein-binding motif. GATA-binding proteins are necessary for the decondensation of heterochromatic regions that hold coding genes, especially in some heteromorphic sex chromosomes that may keep genes related to oocyte development. The (TAA)10 repeat is accumulated in the entire W chromosome, and this microsatellite accumulation is probably involved in the sex chromosome differentiation process and crossover suppression in C. gomesi. These additional data on the W chromosome DNA composition help to explain the evolution of sex chromosomes in Characidium.
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Suwala, Grzegorz, Marie Altmanová, Sofia Mazzoleni, Emmanouela Karameta, Panayiotis Pafilis, Lukáš Kratochvíl, and Michail Rovatsos. "Evolutionary Variability of W-Linked Repetitive Content in Lacertid Lizards." Genes 11, no. 5 (May 11, 2020): 531. http://dx.doi.org/10.3390/genes11050531.
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Lacertid lizards are a widely radiated group of squamate reptiles with long-term stable ZZ/ZW sex chromosomes. Despite their family-wide homology of Z-specific gene content, previous cytogenetic studies revealed significant variability in the size, morphology, and heterochromatin distribution of their W chromosome. However, there is little evidence about the accumulation and distribution of repetitive content on lacertid chromosomes, especially on their W chromosome. In order to expand our knowledge of the evolution of sex chromosome repetitive content, we examined the topology of telomeric and microsatellite motifs that tend to often accumulate on the sex chromosomes of reptiles in the karyotypes of 15 species of lacertids by fluorescence in situ hybridization (FISH). The topology of the above-mentioned motifs was compared to the pattern of heterochromatin distribution, as revealed by C-banding. Our results show that the topologies of the examined motifs on the W chromosome do not seem to follow a strong phylogenetic signal, indicating independent and species-specific accumulations. In addition, the degeneration of the W chromosome can also affect the Z chromosome and potentially also other parts of the genome. Our study provides solid evidence that the repetitive content of the degenerated sex chromosomes is one of the most evolutionary dynamic parts of the genome.
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Odierna, Gaetano, Augusto Gentilli, Marco Zuffi, and Gennaro Aprea. "The karyology of Vipera aspis, V. atra, V. hugyi, and Cerastes vipera." Amphibia-Reptilia 27, no. 1 (2006): 113–19. http://dx.doi.org/10.1163/156853806776052209.
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AbstractIn the current paper we show the results obtained using standard and banding staining methods (Ag-NOR-, CMA3-, C-banding and sequential colorations (or Alu I digestions) + CMA3 + DAPI) in specimens of Cerastes vipera, Vipera aspis, V. atra, and V. hugyi. Cerastes vipera presented chromosomal characters, primitive in snakes, as a karyotype of 2n = 36 chromosomes, with 16 biarmed macrochromosomes and 20 microchromosomes, NORs on one microchromosome pair and absence of cytologically evident sex chromosomes, at least with the methods used. The three taxa of Vipera studied showed chromosomal characters either derived, or primitive or at an initial stage of differentiation. All three species showed a karyotype (derived) of 2n = 42 chromosomes with 22 macro- and 20 micro-chromosomes; they all showed NORs on one micro-chromosome pair and presented Z and W chromosomes at an initial stage of differentiation. Sexchromosomes Z and W, were in fact homomorphic, but the former was near all euchromatic, while the W chromosome was almost completely heterochromatic. All the three taxa of Vipera resulted, however, karyologically diversified, mainly due to the number of macro-chromosomes pairs with a centromeric, CMA3 positive heterochromatin: almost all the pairs in V. aspis, two pairs in V. atra and absent in V. hugyi.
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Fuková, Iva, Petr Nguyen, and Frantiek Marec. "Codling moth cytogenetics: karyotype, chromosomal location of rDNA, and molecular differentiation of sex chromosomes." Genome 48, no. 6 (December 1, 2005): 1083–92. http://dx.doi.org/10.1139/g05-063.
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We performed a detailed karyotype analysis in the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), the key pest of pome fruit in the temperate regions of the world. The codling moth karyotype consisted of 2n = 56 chromosomes of a holokinetic type. The chromosomes were classified into 5 groups according to their sizes: extra large (3 pairs), large (3 pairs), medium (15 pairs), small (5 pairs), and dot-like (2 pairs). In pachytene nuclei of both sexes, a curious NOR (nucleolar organizer region) bivalent was observed. It carried 2 nucleoli, each associated with one end of the bivalent. FISH with an 18S ribosomal DNA probe confirmed the presence of 2 clusters of rRNA genes at the opposite ends of the bivalent. In accordance with this finding, 2 homologous NOR chromosomes were identified in mitotic metaphase, each showing hybridization signals at both ends. In highly polyploid somatic nuclei, females showed a large heterochromatin body, the so-called sex chromatin or W chromatin. The heterochromatin body was absent in male nuclei, indicating a WZ/ZZ (female/male) sex chromosome system. In keeping with the sex chromatin status, pachytene oocytes showed a sex chromosome bivalent (WZ) that was easily discernible by its heterochromatic W thread. To study molecular differentiation of the sex chromosomes, we employed genomic in situ hybridization (GISH) and comparative genomic hybridization (CGH). GISH detected the W chromosome by strong binding of the Cy3-labelled, female-derived DNA probe. With CGH, both the Cy3-labelled female-derived probe and Fluor-X labelled male-derived probe evenly bound to the W chromosome. This suggested that the W chromosome is predominantly composed of repetitive DNA sequences occurring scattered in other chromosomes but accumulated in the W chromosome. The demonstrated ways of W chromosome identification will facilitate the development of genetic sexing strains desirable for pest control using the sterile insect technique.Key words: CGH, codling moth, FISH, GISH, genomic hybridization, heterochromatin, holokinetic chromosomes, karyotype, NOR, rDNA, SIT, sex chromosomes.
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de Souza, Marcelo Santos, Rafael Kretschmer, Suziane Alves Barcellos, Alice Lemos Costa, Marcelo de Bello Cioffi, Edivaldo Herculano Corrêa de Oliveira, Analía Del Valle Garnero, and Ricardo José Gunski. "Repeat Sequence Mapping Shows Different W Chromosome Evolutionary Pathways in Two Caprimulgiformes Families." Birds 1, no. 1 (December 11, 2020): 19–34. http://dx.doi.org/10.3390/birds1010004.
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Although birds belonging to order Caprimulgiformes show extensive karyotype variation, data concerning their genomic organization is still scarce, as most studies have presented only results obtained from conventional staining analyses. Nevertheless, some interesting findings have been observed, such as the W chromosome of the Common Potoo, Nyctibius griseus (2n = 86), which has the same morphology and size of the Z chromosome, a rare feature in Neognathae birds. Hence, we aimed to investigate the process by which the W chromosome of this species was enlarged. For that, we analyzed comparatively the chromosome organization of the Common Potoo and the Scissor-tailed Nightjar, Hydropsalis torquata (2n = 74), which presents the regular differentiated sex chromosomes, by applying C-banding, G-banding and mapping of repetitive DNAs (microsatellite repeats and 18S rDNA). Our results showed an accumulation of constitutive heterochromatin in the W chromosome of both species. However, 9 out of 11 microsatellite sequences hybridized in the large W chromosome in the Common Potoo, while none of them hybridized in the W chromosome of the Scissor-tailed Nightjar. Therefore, we can conclude that the accumulation of microsatellite sequences, and consequent increase in constitutive heterochromatin, was responsible for the enlargement of the W chromosome in the Common Potoo. Based on these results, we conclude that even though these two species belong to the same order, their W chromosomes have gone through different evolutionary histories, with an extra step of accumulation of repetitive sequences in the Common Potoo.
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Odierna, G., T. Caprigilone, L. A. Kupriyanova, and E. Olmo. "Further data on sex chromosomes of Lacertidae and a hypothesis on their evolutionary trend." Amphibia-Reptilia 14, no. 1 (1993): 1–11. http://dx.doi.org/10.1163/156853893x00147.
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AbstractSex chromosomes were studied in eight species of lacertid lizards using C-banding, G-banding and restriction enzyme treatment. All of the species showed female heterogamety. The W chromosome was a microchromosome in Lacerta graeca and Ophisops elegans. Two types of W were found in Lacerta vivipara; in specimens from The Netherlands it was metacentric, whereas in specimens from Russia it was acrocentric or subtelocentric. The W chromosome was homomorphic or nearly homomorphic but completely C-banded and heterochromatic in Lacerta agilis, Podarcis hispanica, Algyroides moreoticus and A. nigropunctatus. In was only possible to find sex chromosomes using the G-banding method in Podarcis sicula. The results obtained, together with data in the literature, suggest that sex chromosomes are likely to be present in all Lacertidae and that their differentiation took place repeatedly and independently in different taxa within the family. A model for sex chromosome evolution in the family, in which the starting point was the heterochromatization of the W chromosome, is proposed.
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Hejníčková, Martina, Petr Koutecký, Pavel Potocký, Irena Provazníková, Anna Voleníková, Martina Dalíková, Sander Visser, František Marec, and Magda Zrzavá. "Absence of W Chromosome in Psychidae Moths and Implications for the Theory of Sex Chromosome Evolution in Lepidoptera." Genes 10, no. 12 (December 5, 2019): 1016. http://dx.doi.org/10.3390/genes10121016.
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Moths and butterflies (Lepidoptera) are the largest group with heterogametic females. Although the ancestral sex chromosome system is probably Z0/ZZ, most lepidopteran species have the W chromosome. When and how the W chromosome arose remains elusive. Existing hypotheses place the W origin either at the common ancestor of Ditrysia and Tischeriidae, or prefer independent origins of W chromosomes in these two groups. Due to their phylogenetic position at the base of Ditrysia, bagworms (Psychidae) play an important role in investigating the W chromosome origin. Therefore, we examined the W chromosome status in three Psychidae species, namely Proutia betulina, Taleporia tubulosa, and Diplodoma laichartingella, using both classical and molecular cytogenetic methods such as sex chromatin assay, comparative genomic hybridization (CGH), and male vs. female genome size comparison by flow cytometry. In females of all three species, no sex chromatin was found, no female-specific chromosome regions were revealed by CGH, and a Z-chromosome univalent was observed in pachytene oocytes. In addition, the genome size of females was significantly smaller than males. Overall, our study provides strong evidence for the absence of the W chromosome in Psychidae, thus supporting the hypothesis of two independent W chromosome origins in Tischeriidae and in advanced Ditrysia.
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Zacharopoulou, A. "Polytene chromosome maps in the Medfly Ceratitis capitata." Genome 33, no. 2 (April 1, 1990): 184–97. http://dx.doi.org/10.1139/g90-030.
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Polytene chromosome maps of the five autosomes from salivary gland cells in Ceratitis capitata are presented, and the more characteristic features of each element are described. The correlation of the polytene elements to miotic chromosomes and linkage groups is established by using various Y-autosome and autosome-autosome translocation lines. Two loci, dp (black pupal case) and w (white pupal case), are mapped to the third and fifth chromosome, respectively. In addition to the polytene maps presented, some extra figures of specific chromosomal regions are given for easier identification of each polytene element.Key words: polytene chromosome maps, Ceratitis capitata.
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Gubb, D., A. Zacharopoulou, Y. Livadaras, P. Gourzi, J. Roote, and C. Savakis. "Recovery of a marked translocation strain that will facilitate the isolation of balancer chromosomes in the Mediterranean fruit fly, Ceratitis capitata." Genome 41, no. 2 (April 1, 1998): 256–65. http://dx.doi.org/10.1139/g98-003.
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The results of two screens for mutations and chromosomal aberrations in Ceratitis capitata are presented. Three dominant mutations were recovered, including Sb, which is associated with a homozygous lethal translocation between the third and fifth chromosomes, T(3;5)Sb, with the fifth chromosome breakpoint adjacent to y. The T(3;5)Sb chromosome is maintained by selecting for Sb in a T(3;5)Sb, w2Sb y2wp/ w2y2wp stock and can be used to distinguish between other chromosomes carrying differential combinations of the recessive markers w2y2wp. The ability to isolate particular marked chromosomes is essential in order to recover an inversion-based balancer chromosome. In addition to the recovery of dominant mutations, gamma -ray induced somatic mosaics of w2 and y2 and zygotic w mosaics were found. The generation of zygotic mosaics following mutagenesis can give mutants with a mosaic germ line that fail to breed true in the first generation. A screen of 22 830 irradiated chromosomes failed to recover variegating alleles of w, although such alleles might be recovered in a larger screen. The high frequency of dominant mutations and the instability at the w locus in our stocks implies a background level of dysgenic activity. These results have implications for the construction and long-term maintenance of genetically modified strains.Key words: dominantly marked translocation, somatic mosaic, variegation, medfly.
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Traut, Walther, Virpi Ahola, David A. S. Smith, Ian J. Gordon, and Richard H. ffrench-Constant. "Karyotypes versus Genomes: The Nymphalid Butterflies Melitaea cinxia, Danaus plexippus, and D. chrysippus." Cytogenetic and Genome Research 153, no. 1 (2017): 46–53. http://dx.doi.org/10.1159/000484032.
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The number of sequenced lepidopteran genomes is increasing rapidly. However, the corresponding assemblies rarely represent whole chromosomes and generally also lack the highly repetitive W sex chromosome. Knowledge of the karyotypes can facilitate genome assembly and further our understanding of sex chromosome evolution in Lepidoptera. Here, we describe the karyotypes of the Glanville fritillary Melitaea cinxia (n = 31), the monarch Danaus plexippus (n = 30), and the African queen D. chrysippus (2n = 60 or 59, depending on the source population). We show by FISH that the telomeres are of the (TTAGG)n type, as found in most insects. M. cinxia and D. plexippus have “conventional” W chromosomes which are heterochromatic in meiotic and somatic cells. In D. chrysippus, the W is inconspicuous. Neither telomeres nor W chromosomes are represented in the published genomes of M. cinxia and D. plexippus. Representation analysis in sequenced female and male D. chrysippus genomes detected an evolutionarily old autosome-Z chromosome fusion in Danaus. Conserved synteny of whole chromosomes, so called “macro synteny”, in Lepidoptera permitted us to identify the chromosomes involved in this fusion. An additional and more recent sex chromosome fusion was found in D. chrysippus by karyotype analysis and classical genetics. In a hybrid population between 2 subspecies, D. c. chrysippus and D. c. dorippus, the W chromosome was fused to an autosome that carries a wing colour locus. Thus, cytogenetics and the present state of genome data complement one another to reveal the evolutionary history of the species.
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Rovatsos, Michail, Martina Johnson Pokorná, and Lukáš Kratochvíl. "Differentiation of Sex Chromosomes and Karyotype Characterisation in the Dragonsnake Xenodermus javanicus (Squamata: Xenodermatidae)." Cytogenetic and Genome Research 147, no. 1 (2015): 48–54. http://dx.doi.org/10.1159/000441646.
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Highly differentiated heteromorphic ZZ/ZW sex chromosomes with a heterochromatic W are a basic principle among advanced snakes of the lineage Colubroidea, while other snake lineages generally lack these characteristics. For the first time, we cytogenetically examined the dragonsnake, Xenodermus javanicus, a member of the family Xenodermatidae, which is phylogenetically nested between snake lineages with and without differentiated sex chromosomes. Although most snakes have a karyotype with a stable chromosomal number of 2n = 36, the dragonsnake has an unusual, derived karyotype with 2n = 32 chromosomes. We found that heteromorphic ZZ/ZW sex chromosomes with a heterochromatic W are present in the dragonsnake, which suggests that the emergence of a highly differentiated W sex chromosome within snakes predates the split of Xenodermatidae and the clade including families Pareatidae, Viperidae, Homalopsidae, Lamprophiidae, Elapidae, and Colubridae. Although accumulations of interstitial telomeric sequences have not been previously reported in snakes, by using FISH with a telomeric probe we discovered them in 6 pairs of autosomes as well as in the W sex chromosome of the dragonsnake. Similarly to advanced snakes, the sex chromosomes of the dragonsnake have a significant accumulation of repeats containing a (GATA)n sequence. The results facilitate the dating of the differentiation of sex chromosomes within snakes back to the split between Xenodermatidae and other advanced snakes, i.e. around 40-75 mya.
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Pucci, Marcela Baer, Viviane Nogaroto, Luiz Antonio Carlos Bertollo, Orlando Moreira-Filho, and Marcelo Ricardo Vicari. "The karyotypes and evolution of ZZ/ZW sex chromosomes in the genus Characidium (Characiformes, Crenuchidae)." Comparative Cytogenetics 12, no. 3 (October 2, 2018): 421–38. http://dx.doi.org/10.3897/compcytogen.v12i3.28736.
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Available data on cytotaxonomy of the genus Characidium Reinhardt, 1867, which contains the greatest number of species in the Characidiinae (Crenuchidae), with 64 species widely distributed throughout the Neotropical region, were summarized and reviewed. Most Characidium species have uniform diploid chromosome number (2n) = 50 and karyotype with 32 metacentric (m) and 18 submetacentric (sm) chromosomes. The maintenance of the 2n and karyotypic formula in Characidium implies that their genomes did not experience large chromosomal rearrangements during species diversification. In contrast, the internal chromosomal organization shows a dynamic differentiation among their genomes. Available data indicated the role of repeated DNA sequences in the chromosomal constitution of the Characidium species, particularly, in sex chromosome differentiation. Karyotypes of the most Characidium species exhibit a heteromorphic ZZ/ZW sex chromosome system. The W chromosome is characterized by high rates of repetitive DNA accumulation, including satellite, microsatellite, and transposable elements (TEs), with a varied degree of diversification among species. In the current review, the main Characidium cytogenetic data are presented, highlighting the major features of its karyotype and sex chromosome evolution. Despite the conserved karyotypic macrostructure with prevalent 2n = 50 chromosomes in Characidium, herein we grouped the main cytogenetic information which led to chromosomal diversification in this Neotropical fish group.
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de Oliveira Furo, Ivanete, Rafael Kretschmer, Michelly S. dos Santos, Carlos A. de Lima Carvalho, Ricardo J. Gunski, Patrícia C. M. O'Brien, Malcolm A. Ferguson-Smith, Marcelo B. Cioffi, and Edivaldo H. C. de Oliveira. "Chromosomal Mapping of Repetitive DNAs in Myiopsitta monachus and Amazona aestiva (Psittaciformes, Psittacidae) with Emphasis on the Sex Chromosomes." Cytogenetic and Genome Research 151, no. 3 (2017): 151–60. http://dx.doi.org/10.1159/000464458.
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Here, for the first time, we describe the karyotype of Myiopsitta monachus (Psittacidae, Arini). We found 2n = 48, corresponding to the lowest diploid number observed in Neotropical Psittaciformes so far, with an uncommonly large W chromosome homomorphic to the Z. In order to better understand the evolution of the sex chromosomes in this species, we applied several molecular cytogenetic approaches, including C-banding, FISH mapping of repetitive DNAs (several microsatellite repeats), and whole-chromosome painting on metaphases of M. monachus. For comparison, another species belonging to the same tribe but with a smaller W chromosome (A. aestiva) was also analyzed. The results show that the constitutive heterochromatin has a very diverse distribution pattern in these species revealing heterochromatic blocks in the centromeric region of all chromosomes and in most of the length of the W chromosome in A. aestiva, while in M. monachus they were found in interstitial and telomeric regions. Concerning the microsatellites, only the sequence (CG)n produced signals on the W chromosome of A. aestiva, in the distal region of both arms. However, in M. monachus, (CAA)n, (CAG)n, and (CG)n probes were accumulated on the W chromosome, and, in addition, the sequence (CAG)n also hybridized to heterochromatic regions in macrochromosomes, as well as in microchromosomes. Based on these results, we suggest that the increase in length of the W chromosome in M. monachus is due to the amplification of repetitive elements, which highlights their significant role in the evolutionary process of sex chromosome differentiation.
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Mäkelä, Jarno, and David Sherratt. "SMC complexes organize the bacterial chromosome by lengthwise compaction." Current Genetics 66, no. 5 (April 16, 2020): 895–99. http://dx.doi.org/10.1007/s00294-020-01076-w.
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Abstract Structural maintenance of chromosomes (SMC) complexes are ancient and conserved molecular machines that organize chromosomes in all domains of life. We propose that the principles of chromosome folding needed to accommodate DNA inside a cell in an accessible form will follow similar principles in prokaryotes and eukaryotes. However, the exact contributions of SMC complexes to bacterial chromosome organization have been elusive. Recently, it was shown that the SMC homolog, MukBEF, organizes and individualizes the Escherichia coli chromosome by forming a filamentous axial core from which DNA loops emanate, similar to the action of condensin in mitotic chromosome formation. MukBEF action, along with its interaction with the partner protein, MatP, also facilitates chromosome individualization by directing opposite chromosome arms (replichores) to different cell halves. This contrasts with the situation in many other bacteria, where SMC complexes organise chromosomes in a way that the opposite replichores are aligned along the long axis of the cell. We highlight the similarities and differences of SMC complex contributions to chromosome organization in bacteria and eukaryotes, and summarize the current mechanistic understanding of the processes.
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Xu, Luohao, Simon Yung Wa Sin, Phil Grayson, Scott V. Edwards, and Timothy B. Sackton. "Evolutionary Dynamics of Sex Chromosomes of Paleognathous Birds." Genome Biology and Evolution 11, no. 8 (July 22, 2019): 2376–90. http://dx.doi.org/10.1093/gbe/evz154.
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Abstract Standard models of sex chromosome evolution propose that recombination suppression leads to the degeneration of the heterogametic chromosome, as is seen for the Y chromosome in mammals and the W chromosome in most birds. Unlike other birds, paleognaths (ratites and tinamous) possess large nondegenerate regions on their sex chromosomes (PARs or pseudoautosomal regions). It remains unclear why these large PARs are retained over >100 Myr, and how this retention impacts the evolution of sex chromosomes within this system. To address this puzzle, we analyzed Z chromosome evolution and gene expression across 12 paleognaths, several of whose genomes have recently been sequenced. We confirm at the genomic level that most paleognaths retain large PARs. As in other birds, we find that all paleognaths have incomplete dosage compensation on the regions of the Z chromosome homologous to degenerated portions of the W (differentiated regions), but we find no evidence for enrichments of male-biased genes in PARs. We find limited evidence for increased evolutionary rates (faster-Z) either across the chromosome or in differentiated regions for most paleognaths with large PARs, but do recover signals of faster-Z evolution in tinamou species with mostly degenerated W chromosomes, similar to the pattern seen in neognaths. Unexpectedly, in some species, PAR-linked genes evolve faster on average than genes on autosomes, suggested by diverse genomic features to be due to reduced efficacy of selection in paleognath PARs. Our analysis shows that paleognath Z chromosomes are atypical at the genomic level, but the evolutionary forces maintaining largely homomorphic sex chromosomes in these species remain elusive.
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Furman, Benjamin L. S., Caroline M. S. Cauret, Martin Knytl, Xue-Ying Song, Tharindu Premachandra, Caleb Ofori-Boateng, Danielle C. Jordan, Marko E. Horb, and Ben J. Evans. "A frog with three sex chromosomes that co-mingle together in nature: Xenopus tropicalis has a degenerate W and a Y that evolved from a Z chromosome." PLOS Genetics 16, no. 11 (November 9, 2020): e1009121. http://dx.doi.org/10.1371/journal.pgen.1009121.
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In many species, sexual differentiation is a vital prelude to reproduction, and disruption of this process can have severe fitness effects, including sterility. It is thus interesting that genetic systems governing sexual differentiation vary among—and even within—species. To understand these systems more, we investigated a rare example of a frog with three sex chromosomes: the Western clawed frog, Xenopus tropicalis. We demonstrate that natural populations from the western and eastern edges of Ghana have a young Y chromosome, and that a male-determining factor on this Y chromosome is in a very similar genomic location as a previously known female-determining factor on the W chromosome. Nucleotide polymorphism of expressed transcripts suggests genetic degeneration on the W chromosome, emergence of a new Y chromosome from an ancestral Z chromosome, and natural co-mingling of the W, Z, and Y chromosomes in the same population. Compared to the rest of the genome, a small sex-associated portion of the sex chromosomes has a 50-fold enrichment of transcripts with male-biased expression during early gonadal differentiation. Additionally, X. tropicalis has sex-differences in the rates and genomic locations of recombination events during gametogenesis that are similar to at least two other Xenopus species, which suggests that sex differences in recombination are genus-wide. These findings are consistent with theoretical expectations associated with recombination suppression on sex chromosomes, demonstrate that several characteristics of old and established sex chromosomes (e.g., nucleotide divergence, sex biased expression) can arise well before sex chromosomes become cytogenetically distinguished, and show how these characteristics can have lingering consequences that are carried forward through sex chromosome turnovers.
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Roco, Álvaro S., Adrián Ruiz-García, and Mónica Bullejos. "Interaction between sex-determining genes from two species: clues from Xenopus hybrids." Philosophical Transactions of the Royal Society B: Biological Sciences 376, no. 1833 (July 26, 2021): 20200104. http://dx.doi.org/10.1098/rstb.2020.0104.
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Hybrids provide an interesting model to study the evolution of sex-determining genes and sex chromosome systems as they offer the opportunity to see how independently evolving sex-determining pathways interact in vivo . In this context, the genus Xenopus represents a stimulating model, since species with non-homologous sex chromosomes and different sex-determining genes have been identified. In addition, the possibility of interspecies breeding is favoured in this group, which arose by alloploidization events, with species ploidy ranging from 2 n = 2 x = 20 in X. tropicalis (the only diploid representative of the genus) to 2 n = 12 x = 108 in X. ruwenzoriensis . To study how two sex-determining genes interact in vivo , X. laevis × X. tropicali s hybrids were produced. Gonadal differentiation in these hybrids revealed that the dm-w gene is dominant over X. tropicalis male-determining sex chromosomes (Y or Z), even though the Y chromosome is dominant in X. tropicalis (Y > W>Z). In the absence of the dm-w gene (the Z chromosome from X. laevis is present), the W chromosome from X. tropicalis is able to trigger ovarian development. Testicular differentiation will take place in the absence of W chromosomes from any of the parental species. The dominance/recessivity relationships between these sex-determining loci in the context of either parental genome remains unknown. This article is part of the theme issue ‘Challenging the paradigm in sex chromosome evolution: empirical and theoretical insights with a focus on vertebrates (Part II)’.
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Giovannotti, Massimo, Paola Nisi Cerioni, Tahar Slimani, Andrea Splendiani, Alessio Paoletti, Adnane Fawzi, Ettore Olmo, and Vincenzo Caputo Barucchi. "Cytogenetic Characterization of a Population of Acanthodactylus lineomaculatus Duméril and Bibron, 1839 (Reptilia, Lacertidae), from Southwestern Morocco and Insights into Sex Chromosome Evolution." Cytogenetic and Genome Research 153, no. 2 (2017): 86–95. http://dx.doi.org/10.1159/000484533.
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Acanthodactylus lineomaculatus is now regarded as an ecotype of A. erythrurus with which it has been recently synonymized. Despite the wide range of A. erythrurus, karyological data for this species are scarce and limited to classical cytogenetic studies carried out in individuals from only 2 locations (central Spain and Spanish enclave of Melilla on the northwestern Mediterranean Moroccan coast). Here, for the first time, we cytogenetically characterized individuals of A. lineomaculatus from the southwestern Moroccan Atlantic coast with the aim to increase the karyological knowledge of this wide-ranging species and to assess if any chromosomal changes can be found in this ecotype in comparison to other populations of this species. The diploid number of the individuals investigated is 2n = 38 which is typical of most lacertids. Active NORs were located telomerically in a medium-small pair of chromosomes, and no inactive NORs were detected. C-banding revealed an intensely heterochromatic W chromosome composed of AT-rich (centromere and long arm telomeric region) and GC-rich (most of the long arm) regions, with extended interstitial telomeric sequences. These telomere-like repeats occupy the GC-rich heterochromatin of the W. The DNA composition of the W represents a trait distinguishing A. lineomaculatus (southwestern Morocco) from A. erythrurus from Spain that possess a DAPI-positive (AT-rich) W chromosome. In conclusion, these results add further evidence to the remarkable karyotype conservation in lacertid lizards, although differences in NOR location and in W chromosome structure among populations could suggest an incipient speciation mediated by chromosome changes in this wide-ranging lizard species.
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Yazdi, Homa Papoli, Willian T. A. F. Silva, and Alexander Suh. "Why Do Some Sex Chromosomes Degenerate More Slowly Than Others? The Odd Case of Ratite Sex Chromosomes." Genes 11, no. 10 (September 30, 2020): 1153. http://dx.doi.org/10.3390/genes11101153.
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The hallmark of sex chromosome evolution is the progressive suppression of recombination which leads to subsequent degeneration of the non-recombining chromosome. In birds, species belonging to the two major clades, Palaeognathae (including tinamous and flightless ratites) and Neognathae (all remaining birds), show distinctive patterns of sex chromosome degeneration. Birds are female heterogametic, in which females have a Z and a W chromosome. In Neognathae, the highly-degenerated W chromosome seems to have followed the expected trajectory of sex chromosome evolution. In contrast, among Palaeognathae, sex chromosomes of ratite birds are largely recombining. The underlying reason for maintenance of recombination between sex chromosomes in ratites is not clear. Degeneration of the W chromosome might have halted or slowed down due to a multitude of reasons ranging from selective processes, such as a less pronounced effect of sexually antagonistic selection, to neutral processes, such as a slower rate of molecular evolution in ratites. The production of genome assemblies and gene expression data for species of Palaeognathae has made it possible, during recent years, to have a closer look at their sex chromosome evolution. Here, we critically evaluate the understanding of the maintenance of recombination in ratites in light of the current data. We conclude by highlighting certain aspects of sex chromosome evolution in ratites that require further research and can potentially increase power for the inference of the unique history of sex chromosome evolution in this lineage of birds.
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Fridolfsson, Anna-Karin, and Hans Ellegren. "Molecular Evolution of the Avian CHD1 Genes on the Z and W Sex Chromosomes." Genetics 155, no. 4 (August 1, 2000): 1903–12. http://dx.doi.org/10.1093/genetics/155.4.1903.
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Abstract Genes shared between the nonrecombining parts of the two types of sex chromosomes offer a potential means to study the molecular evolution of the same gene exposed to different genomic environments. We have analyzed the molecular evolution of the coding sequence of the first pair of genes found to be shared by the avian Z (present in both sexes) and W (female-specific) sex chromosomes, CHD1Z and CHD1W. We show here that these two genes evolve independently but are highly conserved at nucleotide as well as amino acid levels, thus not indicating a female-specific role of the CHD1W gene. From comparisons of sequence data from three avian lineages, the frequency of nonsynonymous substitutions (Ka) was found to be higher for CHD1W (1.55 per 100 sites) than for CHD1Z (0.81), while the opposite was found for synonymous substitutions (Ks, 13.5 vs. 22.7). We argue that the lower effective population size and the absence of recombination on the W chromosome will generally imply that nonsynonymous substitutions accumulate faster on this chromosome than on the Z chromosome. The same should be true for the Y chromosome relative to the X chromosome in XY systems. Our data are compatible with a male-biased mutation rate, manifested by the faster rate of neutral evolution (synonymous substitutions) on the Z chromosome than on the female-specific W chromosome.
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Viana, Ezaz, de Bello Cioffi, Jackson Almeida, and Feldberg. "Evolutionary Insights of the ZW Sex Chromosomes in Snakes: A New Chapter Added by the Amazonian Puffing Snakes of the Genus Spilotes." Genes 10, no. 4 (April 9, 2019): 288. http://dx.doi.org/10.3390/genes10040288.
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Amazonian puffing snakes (Spilotes; Colubridae) are snakes widely distributed in the Neotropical region. However, chromosomal data are scarce in this group and, when available, are only limited to karyotype description using conventional staining. In this paper, we focused on the process of karyotype evolution and trends for sex chromosomes in two Amazonian Puffer Snakes (S. pulllatus and S. sulphureus). We performed an extensive karyotype characterization using conventional and molecular cytogenetic approaches. The karyotype of S. sulphureus (presented here for the first time) exhibits a 2n = 36, similar to that previously described in S. pullatus. Both species have highly differentiated ZZ/ZW sex chromosomes, where the W chromosome is highly heterochromatic in S. pullatus but euchromatic in S. sulphureus. Both W chromosomes are homologous between these species as revealed by cross-species comparative genomic hybridization, even with heterogeneous distributions of several repetitive sequences across their genomes, including on the Z and on the W chromosomes. Our study provides evidence that W chromosomes in these two species have shared ancestry.
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Ta, Thanh Dat, Nomar Espinosa Waminal, Thi Hong Nguyen, Remnyl Joyce Pellerin, and Hyun Hee Kim. "Comparative FISH analysis of Senna tora tandem repeats revealed insights into the chromosome dynamics in Senna." Genes & Genomics 43, no. 3 (March 2021): 237–49. http://dx.doi.org/10.1007/s13258-021-01051-w.
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Abstract Background DNA tandem repeats (TRs) are often abundant and occupy discrete regions in eukaryotic genomes. These TRs often cause or generate chromosomal rearrangements, which, in turn, drive chromosome evolution and speciation. Tracing the chromosomal distribution of TRs could therefore provide insights into the chromosome dynamics and speciation among closely related taxa. The basic chromosome number in the genus Senna is 2n = 28, but dysploid species like Senna tora have also been observed. Objective To understand the dynamics of these TRs and their impact on S. tora dysploidization. Methods We performed a comparative fluorescence in situ hybridization (FISH) analysis among nine closely related Senna species and compared the chromosomal distribution of these repeats from a cytotaxonomic perspective by using the ITS1-5.8S-ITS2 sequence to infer phylogenetic relationships. Results Of the nine S. tora TRs, two did not show any FISH signal whereas seven TRs showed similar and contrasting patterns to other Senna species. StoTR01_86, which was localized in the pericentromeric regions in all S. tora, but not at the nucleolar organizer region (NOR) site, was colocalized at the NOR site in all species except in S. siamea. StoTR02_7_tel was mostly localized at chromosome termini, but some species had an interstitial telomeric repeat in a few chromosomes. StoTR05_180 was distributed in the subtelomeric region in most species and was highly amplified in the pericentromeric region in some species. StoTR06_159 was either absent or colocalized in the NOR site in some species, and StoIGS_463, which was localized at the NOR site in S. tora, was either absent or localized at the subtelomeric or pericentromeric regions in other species. Conclusions These data suggest that TRs play important roles in S. tora dysploidy and suggest the involvement of 45S rDNA intergenic spacers in “carrying” repeats during genome reshuffling.
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Kretschmer, Rafael, Vanusa Lilian Lima, Tiago Marafiga Degrandi, Lucia Vinadé, Adriano Luis Schünemann, Analía Del Valle Garnero, and Ricardo José Gunski. "NOR- bearing as a plesiomorphic characteristic in Mimus saturninus (Passeriformes Mimidae)." Journal of Biotechnology and Biodiversity 5, no. 2 (April 4, 2014): 140–47. http://dx.doi.org/10.20873/jbb.uft.cemaf.v5n2.kretschmer.
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The order Passeriformes is the largest group of species karyotyped among birds, however little is known about the cytogenetic of the Mimidae family, registering only karyology basic data (giemsa staining). The aim of this study was to analyze the chromosomal complement from the species Mimus saturninus by conventional staining and differential chromosome banding. Diploid number and chromosome morphology were determined, as well as the distribution pattern of constitutive heterochromatin (CBG-banding), GTG-banding andAgNOR staining (NORs). The Chalk-browed Mockingbird has 2n=80. The first and fourth pairs are submetacentric and the second, third and fifth are acrocentric. The remaining chromosomes pairs of the complement have telocentric morphology. The Z chromosome is submetacentric and the W is metacentric. CBG-banding showed positive staining in the pericentromeric region of most macrochromosomes and microchromosomes and also at Z chromosome, differently from W chromosome which appeared totally heterochromatic. The GTG-banding was similar to Gallus gallus and in other species which have already been GTG-banded. The NORs were identified in a pair of microchromosomes characterized by presenting a remarkable secondary constriction. This can be considered as a plesiomorphic characteristic for M. saturninus once baseline groups (Paleognathae) also showed a pair of microchromosomes bearing NORs.
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Roco, Álvaro S., Allen W. Olmstead, Sigmund J. Degitz, Tosikazu Amano, Lyle B. Zimmerman, and Mónica Bullejos. "Coexistence of Y, W, and Z sex chromosomes in Xenopus tropicalis." Proceedings of the National Academy of Sciences 112, no. 34 (July 27, 2015): E4752—E4761. http://dx.doi.org/10.1073/pnas.1505291112.
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Homomorphic sex chromosomes and rapid turnover of sex-determining genes can complicate establishing the sex chromosome system operating in a given species. This difficulty exists in Xenopus tropicalis, an anuran quickly becoming a relevant model for genetic, genomic, biochemical, and ecotoxicological research. Despite the recent interest attracted by this species, little is known about its sex chromosome system. Direct evidence that females are the heterogametic sex, as in the related species Xenopus laevis, has yet to be presented. Furthermore, X. laevis’ sex-determining gene, DM-W, does not exist in X. tropicalis, and the sex chromosomes in the two species are not homologous. Here we identify X. tropicalis’ sex chromosome system by integrating data from (i) breeding sex-reversed individuals, (ii) gynogenesis, (iii) triploids, and (iv) crosses among several strains. Our results indicate that at least three different types of sex chromosomes exist: Y, W, and Z, observed in YZ, YW, and ZZ males and in ZW and WW females. Because some combinations of parental sex chromosomes produce unisex offspring and other distorted sex ratios, understanding the sex-determination systems in X. tropicalis is critical for developing this flexible animal model for genetics and ecotoxicology.
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Weber, E. Andreas, and Jörg Grunewald. "Cytotaxonomic differentiation of Wilhelmia equina (Linné, 1747) and Wilhelmia lineata (Meigen, 1804) (Diptera: Simuliidae)." Genome 32, no. 4 (August 1, 1989): 589–95. http://dx.doi.org/10.1139/g89-486.
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In most cases the larvae of Wilhelmia equina and W. lineata cannot be distinguished by using classical morphological features. The morphological characteristics of the salivary gland polytene chromosomes allow one to differentiate clearly between the two species. Characteristic for W. equina are the extended region between the centromere, Ctr (transformed centromere), and the nucleolus organizer, NO, in IS, the definitive position of RB (ring of Balbiani) and bulge in IIS, and the fan-shaped IIIL telomere. The chromosomes of W. lineata are marked by complex chromosomal polymorphisms, the altered position of RB and bulge on IIS and by a strong ectopic pairing of centromeres. The comparison of banding patterns provides several intraspecific polymorphic inversions and interspecific fixed rearrangements for species diagnosis. Partial chromosome maps were established. The comparison of the chromosomal banding pattern of Wilhelmia with that of the Simulium standard reveals a whole-arm interchange between chromosomes I and II in Wilhelmia identical with that in Metomphalus, Prosimulium vernale, a form of P. mixtum, and Metacnephia.Key words: cytotaxonomy, Simuliidae, Wilhelmia equina, Wilhelmia lineata, larvae.
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Ellegren, Hans, and Ariane Carmichael. "Multiple and Independent Cessation of Recombination Between Avian Sex Chromosomes." Genetics 158, no. 1 (May 1, 2001): 325–31. http://dx.doi.org/10.1093/genetics/158.1.325.
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Abstract Birds are characterized by female heterogamety; females carry the Z and W sex chromosomes, while males have two copies of the Z chromosome. We suggest here that full differentiation of the Z and W sex chromosomes of birds did not take place until after the split of major contemporary lineages, in the late Cretaceous. The ATP synthase α-subunit gene is now present in one copy each on the nonrecombining part of the W chromosome (ATP5A1W) and on the Z chromosome (ATP5A1Z). This gene seems to have evolved on several independent occasions, in different lineages, from a state of free recombination into two sex-specific and nonrecombining variants. ATP5A1W and ATP5A1Z are thus more similar within orders, relative to what W (or Z) are between orders. Moreover, this cessation of recombination apparently took place at different times in different lineages (estimated at 13, 40, and 65 million years ago in Ciconiiformes, Galliformes, and Anseriformes, respectively). We argue that these observations are the result of recent and traceable steps in the process where sex chromosomes gradually cease to recombine and become differentiated. Our data demonstrate that this process, once initiated, may occur independently in parallel in sister lineages.
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Konrath, Alicia, Ann-Kathrin Schmidt, and Holger Bastians. "DNA-Replikationsstress, Mitose und genomische Instabilität." BIOspektrum 27, no. 1 (February 2021): 10–13. http://dx.doi.org/10.1007/s12268-021-1525-5.
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AbstractChromosomal instability (CIN) is a hallmark of cancer and contributes to tumorigenesis and tumor progression. While structural CIN (S-CIN) leads to structural chromosome aberrations, whole chromosome instability (W-CIN) is defined by perpetual gains or losses of chromosomes during mitosis causing aneuploidy. Mitotic defects, but also abnormal DNA replication (replication stress) can lead to W-CIN. However, the functional link between replication stress, mitosis and aneuploidy is little understood.
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Rogers, Thea F., Tommaso Pizzari, and Alison E. Wright. "Multi-Copy Gene Family Evolution on the Avian W Chromosome." Journal of Heredity 112, no. 3 (March 24, 2021): 250–59. http://dx.doi.org/10.1093/jhered/esab016.
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Abstract The sex chromosomes often follow unusual evolutionary trajectories. In particular, the sex-limited chromosomes frequently exhibit a small but unusual gene content in numerous species, where many genes have undergone massive gene amplification. The reasons for this remain elusive with a number of recent studies implicating meiotic drive, sperm competition, genetic drift, and gene conversion in the expansion of gene families. However, our understanding is primarily based on Y chromosome studies as few studies have systematically tested for copy number variation on W chromosomes. Here, we conduct a comprehensive investigation into the abundance, variability, and evolution of ampliconic genes on the avian W. First, we quantified gene copy number and variability across the duck W chromosome. We find a limited number of gene families as well as conservation in W-linked gene copy number across duck breeds, indicating that gene amplification may not be such a general feature of sex chromosome evolution as Y studies would initially suggest. Next, we investigated the evolution of HINTW, a prominent ampliconic gene family hypothesized to play a role in female reproduction and oogenesis. In particular, we investigated the factors driving the expansion of HINTW using contrasts between modern chicken and duck breeds selected for different female-specific selection regimes and their wild ancestors. Although we find the potential for selection related to fecundity in explaining small-scale gene amplification of HINTW in the chicken, purifying selection seems to be the dominant mode of evolution in the duck. Together, this challenges the assumption that HINTW is key for female fecundity across the avian phylogeny.
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Küpper, Clemens, Jakob Augustin, Scott Edwards, Tamás Székely, András Kosztolányi, Terry Burke, and Daniel E. Janes. "Triploid plover female provides support for a role of the W chromosome in avian sex determination." Biology Letters 8, no. 5 (May 30, 2012): 787–89. http://dx.doi.org/10.1098/rsbl.2012.0329.
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Two models, Z Dosage and Dominant W , have been proposed to explain sex determination in birds, in which males are characterized by the presence of two Z chromosomes, and females are hemizygous with a Z and a W chromosome. According to the Z Dosage model, high dosage of a Z-linked gene triggers male development, whereas the Dominant W model postulates that a still unknown W-linked gene triggers female development. Using 33 polymorphic microsatellite markers, we describe a female triploid Kentish plover Charadrius alexandrinus identified by characteristic triallelic genotypes at 14 autosomal markers that produced viable diploid offspring. Chromatogram analysis showed that the sex chromosome composition of this female was ZZW. Together with two previously described ZZW female birds, our results suggest a prominent role for a female determining gene on the W chromosome. These results imply that avian sex determination is more dynamic and complex than currently envisioned.
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Lisachov, Artem, Daria Andreyushkova, Guzel Davletshina, Dmitry Prokopov, Svetlana Romanenko, Svetlana Galkina, Alsu Saifitdinova, Evgeniy Simonov, Pavel Borodin, and Vladimir Trifonov. "Amplified Fragments of an Autosome-Borne Gene Constitute a Significant Component of the W Sex Chromosome of Eremias velox (Reptilia, Lacertidae)." Genes 12, no. 5 (May 20, 2021): 779. http://dx.doi.org/10.3390/genes12050779.
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Heteromorphic W and Y sex chromosomes often experience gene loss and heterochromatinization, which is frequently viewed as their “degeneration”. However, the evolutionary trajectories of the heterochromosomes are in fact more complex since they may not only lose but also acquire new sequences. Previously, we found that the heterochromatic W chromosome of a lizard Eremias velox (Lacertidae) is decondensed and thus transcriptionally active during the lampbrush stage. To determine possible sources of this transcription, we sequenced DNA from a microdissected W chromosome sample and a total female DNA sample and analyzed the results of reference-based and de novo assembly. We found a new repetitive sequence, consisting of fragments of an autosomal protein-coding gene ATF7IP2, several SINE elements, and sequences of unknown origin. This repetitive element is distributed across the whole length of the W chromosome, except the centromeric region. Since it retained only 3 out of 10 original ATF7IP2 exons, it remains unclear whether it is able to produce a protein product. Subsequent studies are required to test the presence of this element in other species of Lacertidae and possible functionality. Our results provide further evidence for the view of W and Y chromosomes as not just “degraded” copies of Z and X chromosomes but independent genomic segments in which novel genetic elements may arise.
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Green, David M. "Muller's Ratchet and the evolution of supernumerary chromosomes." Genome 33, no. 6 (December 1, 1990): 818–24. http://dx.doi.org/10.1139/g90-123.
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Supernumerary chromosomes arise from portions of the normal chromosome complement through nondisjunction, fragmentation, or other mechanisms. Once present in the genome, they are subject to virtually the same genetic conditions that affect the evolutionary degeneration of heteromorphic sex chromosomes. Y or W chromosomes occur only in the presence of X or Z chromosomes, respectively, just as supernumeraries never occur except in the presence of the complete regular karyotype containing their progenitor sequences. Thus, mechanisms that can account for the evolution of sex-chromosome heteromorphism can also be invoked to explain the degeneration process of supernumerary chromosomes after their origination. Incipient supernumeraries initially have genes identical with those on progenitor chromosomes. This frees them from the evolutionary constraint of carrying nonduplicated genetic information, just as in Y and W chromosomes during early stages of sex-chromosome differentiation. The degeneration of supernumerary chromosomes may thus proceed via the mechanism of Muller's Ratchet. This hypothesis predicts that supernumerary chromosomes should lose functional loci, lose sequence homology with the regular genome, and gain heterochromatin over time, resulting in multiple heteromorphic forms of degenerate supernumeraries within and between populations, as is commonly observed.Key words: supernumerary chromosomes, B chromosomes, evolution, origin, Muller's Ratchet.
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Leclercq, Sébastien, Julien Thézé, Mohamed Amine Chebbi, Isabelle Giraud, Bouziane Moumen, Lise Ernenwein, Pierre Grève, Clément Gilbert, and Richard Cordaux. "Birth of a W sex chromosome by horizontal transfer of Wolbachia bacterial symbiont genome." Proceedings of the National Academy of Sciences 113, no. 52 (December 6, 2016): 15036–41. http://dx.doi.org/10.1073/pnas.1608979113.
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Sex determination is a fundamental developmental pathway governing male and female differentiation, with profound implications for morphology, reproductive strategies, and behavior. In animals, sex differences between males and females are generally determined by genetic factors carried by sex chromosomes. Sex chromosomes are remarkably variable in origin and can differ even between closely related species, indicating that transitions occur frequently and independently in different groups of organisms. The evolutionary causes underlying sex chromosome turnover are poorly understood, however. Here we provide evidence indicating that Wolbachia bacterial endosymbionts triggered the evolution of new sex chromosomes in the common pillbug Armadillidium vulgare. We identified a 3-Mb insert of a feminizing Wolbachia genome that was recently transferred into the pillbug nuclear genome. The Wolbachia insert shows perfect linkage to the female sex, occurs in a male genetic background (i.e., lacking the ancestral W female sex chromosome), and is hemizygous. Our results support the conclusion that the Wolbachia insert is now acting as a female sex-determining region in pillbugs, and that the chromosome carrying the insert is a new W sex chromosome. Thus, bacteria-to-animal horizontal genome transfer represents a remarkable mechanism underpinning the birth of sex chromosomes. We conclude that sex ratio distorters, such as Wolbachia endosymbionts, can be powerful agents of evolutionary transitions in sex determination systems in animals.
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Gunski, Ricardo J., Rafael Kretschmer, Marcelo Santos de Souza, Ivanete de Oliveira Furo, Suziane A. Barcellos, Alice L. Costa, Marcelo B. Cioffi, Edivaldo H. C. de Oliveira, and Analía del Valle Garnero. "Evolution of Bird Sex Chromosomes Narrated by Repetitive Sequences: Unusual W Chromosome Enlargement in Gallinula melanops (Aves: Gruiformes: Rallidae)." Cytogenetic and Genome Research 158, no. 3 (2019): 152–59. http://dx.doi.org/10.1159/000501381.
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Among birds, species with the ZZ/ZW sex determination system generally show significant differences in morphology and size between the Z and W chromosomes (with the W usually being smaller than the Z). In the present study, we report for the first time the karyotype of the spot-flanked gallinule (Gallinula melanops) by means of classical and molecular cytogenetics. The spot-flanked gallinule has 2n = 80 (11 pairs of macrochromosomes and 29 pairs of microchromosomes) with an unusual W chromosome that is larger than the Z. Besides being totally heterochromatic, it has a secondary constriction in its long arm corresponding to the nucleolar organizer region, as confirmed by both silver staining and mapping of 18S rDNA probes. This is an unprecedented fact among birds. Additionally, 18S rDNA sites were also observed in 6 microchromosomes, while 5S rDNA was found in just 1 microchromosomal pair. Seven out of the 11 used microsatellite sequences were found to be accumulated in microchromosomes, and 6 microsatellite sequences were found in the W chromosome. In addition to the involvement of heterochromatin and repetitive DNAs in the differentiation of the large W chromosome, the results also show an alternative scenario that highlights the plasticity that shapes the evolutionary history of bird sex chromosomes.
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Kretschmer, Rafael, Ricardo José Gunski, Analía del Valle Garnero, Thales Renato Ochotorena de Freitas, Gustavo Akira Toma, Marcelo de Bello Cioffi, Edivaldo Herculano Corrêa de Oliveira, Rebecca E. O’Connor, and Darren K. Griffin. "Chromosomal Analysis in Crotophaga ani (Aves, Cuculiformes) Reveals Extensive Genomic Reorganization and an Unusual Z-Autosome Robertsonian Translocation." Cells 10, no. 1 (December 22, 2020): 4. http://dx.doi.org/10.3390/cells10010004.
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Although cytogenetics studies in cuckoos (Aves, Cuculiformes) have demonstrated an interesting karyotype variation, such as variations in the chromosome morphology and diploid number, their chromosome organization and evolution, and relation with other birds are poorly understood. Hence, we combined conventional and molecular cytogenetic approaches to investigate chromosome homologies between chicken and the smooth-billed ani (Crotophaga ani). Our results demonstrate extensive chromosome reorganization in C. ani, with interchromosomal rearrangements involving macro and microchromosomes. Intrachromosomal rearrangements were observed in some macrochromosomes, including the Z chromosome. The most evolutionary notable finding was a Robertsonian translocation between the microchromosome 17 and the Z chromosome, a rare event in birds. Additionally, the simple short repeats (SSRs) tested here were preferentially accumulated in the microchromosomes and in the Z and W chromosomes, showing no relationship with the constitutive heterochromatin regions, except in the W chromosome. Taken together, our results suggest that the avian sex chromosome is more complex than previously postulated and revealed the role of microchromosomes in the avian sex chromosome evolution, especially cuckoos.
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Sigeman, Hanna, Suvi Ponnikas, Elin Videvall, Hongkai Zhang, Pallavi Chauhan, Sara Naurin, and Bengt Hansson. "Insights into Avian Incomplete Dosage Compensation: Sex-Biased Gene Expression Coevolves with Sex Chromosome Degeneration in the Common Whitethroat." Genes 9, no. 8 (July 26, 2018): 373. http://dx.doi.org/10.3390/genes9080373.
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Non-recombining sex chromosomes (Y and W) accumulate deleterious mutations and degenerate. This poses a problem for the heterogametic sex (XY males; ZW females) because a single functional gene copy often implies less gene expression and a potential imbalance of crucial expression networks. Mammals counteract this by dosage compensation, resulting in equal sex chromosome expression in males and females, whereas birds show incomplete dosage compensation with significantly lower expression in females (ZW). Here, we study the evolution of Z and W sequence divergence and sex-specific gene expression in the common whitethroat (Sylvia communis), a species within the Sylvioidea clade where a neo-sex chromosome has been formed by a fusion between an autosome and the ancestral sex chromosome. In line with data from other birds, females had lower expression than males at the majority of sex-linked genes. Results from the neo-sex chromosome region showed that W gametologs have diverged functionally to a higher extent than their Z counterparts, and that the female-to-male expression ratio correlated negatively with the degree of functional divergence of these gametologs. We find it most likely that sex-linked genes are being suppressed in females as a response to W chromosome degradation, rather than that these genes experience relaxed selection, and thus diverge more, by having low female expression. Overall, our data of this unique avian neo-sex chromosome system suggest that incomplete dosage compensation evolves, at least partly, through gradual accumulation of deleterious mutations at the W chromosome and declining female gene expression.
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Sundström, Hannah, Matthew T. Webster, and Hans Ellegren. "Is the Rate of Insertion and Deletion Mutation Male Biased?: Molecular Evolutionary Analysis of Avian and Primate Sex Chromosome Sequences." Genetics 164, no. 1 (May 1, 2003): 259–68. http://dx.doi.org/10.1093/genetics/164.1.259.
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Abstract The rate of mutation for nucleotide substitution is generally higher among males than among females, likely owing to the larger number of DNA replications in spermatogenesis than in oogenesis. For insertion and deletion (indel) mutations, data from a few human genetic disease loci indicate that the two sexes may mutate at similar rates, possibly because such mutations arise in connection with meiotic crossing over. To address origin- and sex-specific rates of indel mutation we have conducted the first large-scale molecular evolutionary analysis of indels in noncoding DNA sequences from sex chromosomes. The rates are similar on the X and Y chromosomes of primates but about twice as high on the avian Z chromosome as on the W chromosome. The fact that indels are not uncommon on the nonrecombining Y and W chromosomes excludes meiotic crossing over as the main cause of indel mutation. On the other hand, the similar rates on X and Y indicate that the number of DNA replications (higher for Y than for X) is also not the main factor. Our observations are therefore consistent with a role of both DNA replication and recombination in the generation of short insertion and deletion mutations. A significant excess of deletion compared to insertion events is observed on the avian W chromosome, consistent with gradual DNA loss on a nonrecombining chromosome.
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Peona, Valentina, Octavio M. Palacios-Gimenez, Julie Blommaert, Jing Liu, Tri Haryoko, Knud A. Jønsson, Martin Irestedt, Qi Zhou, Patric Jern, and Alexander Suh. "The avian W chromosome is a refugium for endogenous retroviruses with likely effects on female-biased mutational load and genetic incompatibilities." Philosophical Transactions of the Royal Society B: Biological Sciences 376, no. 1833 (July 26, 2021): 20200186. http://dx.doi.org/10.1098/rstb.2020.0186.
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It is a broadly observed pattern that the non-recombining regions of sex-limited chromosomes (Y and W) accumulate more repeats than the rest of the genome, even in species like birds with a low genome-wide repeat content. Here, we show that in birds with highly heteromorphic sex chromosomes, the W chromosome has a transposable element (TE) density of greater than 55% compared to the genome-wide density of less than 10%, and contains over half of all full-length (thus potentially active) endogenous retroviruses (ERVs) of the entire genome. Using RNA-seq and protein mass spectrometry data, we were able to detect signatures of female-specific ERV expression. We hypothesize that the avian W chromosome acts as a refugium for active ERVs, probably leading to female-biased mutational load that may influence female physiology similar to the ‘toxic-Y’ effect in Drosophila males. Furthermore, Haldane's rule predicts that the heterogametic sex has reduced fertility in hybrids. We propose that the excess of W-linked active ERVs over the rest of the genome may be an additional explanatory variable for Haldane's rule, with consequences for genetic incompatibilities between species through TE/repressor mismatches in hybrids. Together, our results suggest that the sequence content of female-specific W chromosomes can have effects far beyond sex determination and gene dosage. This article is part of the theme issue ‘Challenging the paradigm in sex chromosome evolution: empirical and theoretical insights with a focus on vertebrates (Part II)’.
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Ezaz, Tariq, and Janine E. Deakin. "Repetitive Sequence and Sex Chromosome Evolution in Vertebrates." Advances in Evolutionary Biology 2014 (September 11, 2014): 1–9. http://dx.doi.org/10.1155/2014/104683.
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Sex chromosomes are the most dynamic entity in any genome having unique morphology, gene content, and evolution. They have evolved multiple times and independently throughout vertebrate evolution. One of the major genomic changes that pertain to sex chromosomes involves the amplification of common repeats. It is hypothesized that such amplification of repeats facilitates the suppression of recombination, leading to the evolution of heteromorphic sex chromosomes through genetic degradation of Y or W chromosomes. Although contrasting evidence is available, it is clear that amplification of simple repetitive sequences played a major role in the evolution of Y and W chromosomes in vertebrates. In this review, we present a brief overview of the repetitive DNA classes that accumulated during sex chromosome evolution, mainly focusing on vertebrates, and discuss their possible role and potential function in this process.
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Itoh, Yuichiro, Kathy Kampf, and Arthur P. Arnold. "Molecular cloning of zebra finch W chromosome repetitive sequences: evolution of the avian W chromosome." Chromosoma 117, no. 2 (October 31, 2007): 111–21. http://dx.doi.org/10.1007/s00412-007-0130-8.
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Rojek, Aleksandra, Karolina Kwasiuk, Monika Obara-Moszyńska, Zofia Kolesińska, and Marek Niedziela. "Y chromosome in Turner syndrome." Pediatric Endocrinology Diabetes and Metabolism 23, no. 1 (2017): 37–41. http://dx.doi.org/10.18544/pedm-23.01.0072.
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